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Lecanora helmutii , a New Species from the Lecanora symmicta Group from
Tasmania

Article  in  Herzogia · October 2018

DOI: 10.13158/heia.31.1.2018.639

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Herzogia 31 (1) Teil 2, 2018: 639 –649 639

Lecanora helmutii, a new species from the

Lecanora symmicta group from Tasmania

Sergio Pérez-Ortega* & Gintaras Kantvilas

Abstract: Pérez-Ortega S. & Kantvilas G. 2018: Lecanora helmutii, a new species from the Lecanora symmicta
group from Tasmania. – Herzogia 31: 639 –649.
Lecanora helmutii Pérez-Ortega & Kantvilas from Tasmania is described as new to science. It shows characters typi-
cal of the Lecanora symmicta group, such as biatorine apothecia, Lecanora-type asci, and simple, hyaline ascospores.
It is further characterized by whitish, discrete soralia and the presence of usnic acid and zeorin as major compounds.
Using the universal fungal barcode nrITS and Bayesian and maximum likelihood inference of the phylogenetic rela-
tionships, we confirm its placement in the Lecanora symmicta group. Lecanora strobilina and L. strobilinoides are
sequenced for the first time and their affiliation to the Lecanora symmicta group is likewise confirmed.

Zusammenfassung: Pérez-Ortega S. & Kantvilas G. 2018: Lecanora helmutii, eine neue Art aus der Lecanora
symmicta-Gruppe aus Tasmanien. – Herzogia 31: 639 –649.
Lecanora helmutii Pérez-Ortega & Kantvilas aus Tasmanien wird neu beschrieben. Die Art zeigt für die Lecanora
symmicta-Gruppe typische Merkmale, wie biatorine Apothecien, Asci vom Lecanora-Typ und einfache, hyaline
Ascosporen. Weitere Merkmale sind die weißlichen, abgegrenzten Sorale und das Auftreten von Usninsäure und Zeorin
als Hauptinhaltsstoffe. Mithilfe des universellen Pilz-Barcodes nrITS und mittels Bayesianischer und Maximum-
Likelihood-Methoden zur Analyse phylogenetischer Verwandtschaftsverhältnisse konnten wir ihre Zugehörigkeit zur
Lecanora symmicta-Gruppe zeigen. Erstmals werden Sequenzdaten von Lecanora strobilina und L. strobilinoides
vorgelegt und damit ebenso deren verwandtschaftliche Nähe zur Lecanora symmicta-Gruppe bestätigt.

Key words: Australia, Lecanoraceae, lichen-forming fungi, zeorin, usnic acid.

Introduction
The genus Lecanora Ach. in a broad sense is defined by crustose thalli, apothecia with margins
containing algal cells, and Lecanora-type asci with simple, colourless ascospores (Ryan et al.
2004). With more than 500 species, it is the second richest genus of lichenized fungi (Lücking
et al. 2016). Several recent molecular studies have suggested that this broad, morphological
concept of the genus is artificial, and demonstrated the existence of numerous monophyletic
clades (Arup & Grube 2000, Grube et al. 2004, Papong et al. 2013, Pérez-Ortega et
al. 2010, Zhao et al. 2016), many of which have been recognized as different genera in the
past (Choisy 1929, Eigler 1969). This has led to new taxonomic arrangements in the genus
(Rodríguez Flakus & Printzen 2014, Zhao et al. 2016).
One of these well-defined and supported groups corresponds to the Lecanora symmicta group
(Pérez-Ortega et al. 2010). Species of this group have previously been included within the

* Corresponding author
640 Herzogia 31 (1) Teil 2, 2018

Lecanora varia group (= Straminella M.Choisy) on account of their yellowish thallus that
contains usnic acid (Śliwa & Wetmore 2000), a character that also occurs in Lecanora s. str.
(Lumbsch et al. 1995). The L. symmicta group can be defined by the exclusion of the thalline
margin in the apothecia (at least in mature specimens), giving them a typical biatorine ap-
pearance, the Lecanora-type ascus (but see Rodriguez Flakus & Printzen 2014), and the
presence of usnic acid (Guzow-Krzemińska et al. 2017, Printzen 2001, Printzen & May
2002, Rodriguez Flakus & Printzen 2014). Relationships between the L. symmicta group
and other clades in Lecanora s. lat. are not resolved (Pérez-Ortega et al. 2010). Based on
five loci, Rodriguez Flakus & Printzen (2014) demonstrated its relationship with the re-
cently described genus Palicella Flakus & Printzen and with Pyrrhospora Körb. On the other
hand, Zhao et al. (2016), using a 6-locus data set, placed it in a well-supported clade together
with members of Protoparmeliopsis M.Choisy, Myriolecis Clem., Rhizoplaca Zopf and the L.
polytropa group.
The L. symmicta group has not been thoroughly revised and remains poorly known in the
Southern Hemisphere. Lecanora symmicta (Ach.) Ach. itself has been reported from Australia
(Lumbsch & Elix 2004) although it is not clear whether this corresponds exactly to the
Northern Hemisphere taxon since there are several additional taxa close to the concept of L.
symmicta that occur in the region, some of which remain undescribed (Kantvilas & LaGreca
2008, Kantvilas 2012). During studies of the corticolous lichen biota of coastal woodlands in
Tasmania, the second author collected a lichen resembling members of the Lecanora symmicta
group and containing usnic acid and zeorin as major compounds. This taxon also had discrete,
whitish soralia, a character rarely seen in the group. Here we present our investigation of this
lichen and describe it as new to science. We carried out molecular analyses of the barcoding
ITS region in order to establish its phylogenetic affinities and to confirm its placement in the L.
symmicta group. We also include other putative members of the group that have not been ana-
lyzed previously: L. strobilina (Spreng.) Kieff. and L. strobilinoides Giralt & Gómez-Bolea.

Material and methods

Molecular analyses
Sequence alignment and phylogenetic analysis
Slices of the apothecia were used for DNA isolation following inspection under the compound
microscope to check for algal or fungal contaminants. Genomic DNA was isolated using a
CTAB-based method (Werth et al. 2016). We amplified the nrITS using the primers pairs
ITS1F-Kyo2 and ITS4-Kyo2 (Toju et al. 2012). PCR reactions were performed in a total
volume of 15 µl, containing 2 µl of template DNA, 0.5 µl of each primer (10 µm), 6.5 µl of
MyTaq Mix (MyTaq DNA Polymerase [Bioline] and dNTPs); distilled water was added to
reach the final volume. PCR amplifications were performed in an Eppendorf Mastercycler
EPgradient S thermal cycler. PCR conditions were: an initial 4 min heating phase at 95ºC,
30 more cycles of 1 min at 94ºC, 1:15 min at 52ºC and 1:30 min at 94ºC, and a final extension
step of 7 min at 72ºC. PCR products were purified using ExoSAP-IT™ PCR Product Cleanup
Reagent (ThermoFisher) according to manufacturer instructions, and sequenced by Macrogen
Inc. (Madrid, Spain) using the same primer set as for PCR amplification. Sequence contigs
were assembled using SeqMan v.12.0 (Lasergene, DNA Star Inc., WI, USA).
Sequences obtained in our study (Table 1) were aligned with members of the L. symmicta
group following Guzow-Krzemińska et al. (2017) using MAFFT v. 7.0. (Katoh et al. 2009),
Pérez-Ortega & Kantvilas: Lecanora helmutii, a new species from Tasmania 641

as implemented in GeneiousTM v. 9.0.4. (Biomatters Ltd, New Zealand) using default param-
eters, and subsequently inspected by eye. Palicella schizochromatica (Pérez-Ortega, T.Sprib.
& Printzen) Rodr.Flakus & Printzen was used as outgroup. Gblocks 0.91b (Castresana 2000)
implemented at http://molevol.cmima.csic.es/castresana/Gblocks_server.html, was used to re-
move ambiguously aligned regions and large gaps using the least stringent parameters, but
allowing gaps in 50 % of the sequences. nrITS was subdivided in ITS1, 5.8S and ITS2 re-
gions and optimal nucleotide substitution models for the Bayesian analysis for each genomic
region were established by means of jModelTest v.2 (Darriba et al. 2012) using the Akaike
Information Criterion (ITS1 & ITS2:GTR+G; 5.8S: JC).
IQ-TREE v.1.6. (Kalyaanamoorthy et al. 2017, Nguyen et al. 2015), implemented online
at http://iqtree.cibiv.univie.ac.at, was used for the heuristic search of the maximum likelihood
(ML) tree and the simultaneous inference of the substitution model for each data partition and
the optimal partitioning scheme. Branch support was estimated with the ultrafast bootstrap
algorithm (Minh et al. 2013).
Bayesian inference of the phylogenetic relationships was carried out in MrBayes v.3.2
(Ronquist et al. 2012). Settings included two parallel runs with four chains over 30 M gen-
erations sampling every 1000th step. The first 25 % of saved data was discarded as ‘burn-in’.
Convergence of chains was confirmed by the convergent diagnostic of the Potential Scale
Reduction Factor (PSRF), which approached 1 and inspected visually in Tracer v. 1.6 (http://
tree.bio.ed.ac.uk/software/tracer/). Phylogenetic trees were visualized in FigTree v. 1.4.1
(http://tree.bio.ed.ac.uk/software/figtree/) and Adobe Illustrator CS2® was used for artwork.
Strongly supported nodes were defined as those with ultrafast bootstrap value ≥ 95 % and
Bayesian posterior probability ≥ 0.95, shown in bold in figure 1.

Table 1: Collection data, herbarium numbers and GenBank accession numbers from the newly sequenced Lecanora
specimens in this study.

Country Collector Herbarium Laborat. nrITS accession

number code number
Lecanora helmutii Australia Kantvilas 460/11 MA-Lichen S844 MG973240
(ISOTYPE) 19506
Lecanora strobilina Portugal S. Pérez-Ortega 5745 MA-Lichen S4107 MG973237
& M. Blázquez 19509
Lecanora strobilina Spain S. Pérez-Ortega 5270 MA-Lichen S4112 MG973235
& S. Prats 19510
Lecanora strobilina Spain S. Pérez-Ortega 5267 MA-Lichen S4114 MG973236
& S. Prats 19511
Lecanora strobilinoides Spain I. Garrido-Benavent sn MA-Lichen S3499 MG973239
19508
Lecanora strobilinoides Spain I. Garrido-Benavent sn MA-Lichen S3500 MG973238
19507

Morphological analyses
Specimens were examined under a Leica S8APO dissecting microscope. Habit photographs
were taken using a Canon 70D coupled with a Canon MP-E 65 mm f/2.8 1-5x macro lens.
Hand-cut sections of apothecia and thallus were observed using either a Zeiss Axioplan 2 fitted
with Nomarski differential interference contrast (DIC) or a Nikon Eclipse E200 fitted with a
set for polarized light. Photographs were taken with a Zeiss Axiovision 4.8 image analyser sys-
642 Herzogia 31 (1) Teil 2, 2018

Figure 1: nrITS Bayesian phylogenetic tree of the Lecanora symmicta group obtained in MrBayes (ML analysis
recovered the same topology). Bold branches depict both ultrafast bootstrap (B) and posterior probability (PP) sup-
port; figures are given above or below branches (B/PP). Lecanora helmutii, L. strobilina and L. strobilinoides are
highlighted in light grey.

tem. Colour reactions were observed using 50 % HNO3 (N), 10 % KOH (K) and Lugol’s iodine
solution (I), the latter both with (K/I) and without pretreatment with K. The measurements of
ascospores are presented as minimum–(average ±standard deviation)–maximum values fol-
lowed by the number of measurements (n). Thin layer chromatography analyses were carried
out following standard methods (Orange et al. 2001).

Results
Molecular analyses
We analyzed 39 nrITS sequences, six of them newly generated. Alignment was 545 bp long,
424 of which remained after Gblocks removal of ambiguously aligned regions and gaps.
Pérez-Ortega & Kantvilas: Lecanora helmutii, a new species from Tasmania 643

Bayesian and maximum likelihood analyses retrieved 50 % majority rule and single best trees
respectively with similar topologies, so only the tree recovered using Bayesian inference
is shown in figure 1. Overall supported relationships among the species are rather similar
to those inferred by Guzow-Krzemińska et al. (2017) and will not be discussed in detail.
The inclusion of the new taxon in the L. symmicta group is well supported by both analyses.
However, its position it is not clear, appearing as basal to the core of the group in the Bayesian
analysis and basal to the clade comprising L. aitema (Ach.) Hepp and Lecanora flavoleprosa
Tønsberg in the ML analysis, although neither of those relationships is statistically supported.
The two specimens of Lecanora strobilinoides Giralt & Gómez-Bolea included in the analyses
appear as members of the L. symmicta group, sister to L. fulvastra Kremp. On the other hand,
specimens of L. strobilina (Spreng.) Kieff. appear in an unresolved clade together with speci-
mens of L. expallens Ach.

Taxonomy
Lecanora helmutii Pérez-Ortega & Kantvilas, species nova (figure 2)
MycoBank number: MB 824355
Diagnosis: Characterized by a sorediate thallus containing usnic acid and zeorin as major compounds,
with the soralia initially discrete, 0.2– 0.5 mm wide, biatorine, lemon yellowish to cream-coloured,
lightly pruinose apothecia, Lecanora-type asci and simple, hyaline ascospores,10 –15 × 4 – 6.5 µm.
Type: Australia, Tasmania, eastern side of Stanley Highway, 40°47'S, 145°16'E, 5 m alt., on Banksia
marginata in Melaleuca-dominated coastal swampy woodland. Kantvilas 460/11, 28 November 2011
[holotype in HO (563771); isotype in MA-Lichen (19506)].
Etymology: The new species is named after Dr. Helmut Mayrhofer (Graz, Austria), a highly respected
lichenologist and friend and colleague to us both.
Description: Thallus effuse, ecorticate, not well delimited, whitish, sorediate; soralia discrete, arising
in groups, becoming confluent in parts of the thallus, ± roundish, excavate to erumpent, c. 0.2– 0.5 mm
wide; soredia to 25 µm diam; medulla I–. Photobiont Trebouxia-like, with ± globose cells to 20 µm diam.
Apothecia biatorine, to 0.5 mm diam, scattered to crowded, irregularly roundish, occurring on both
sorediate and esorediate areas of the thallus; disc lemon yellowish to cream-coloured, delicately prui-
nose, convex or, more rarely, plane; excipulum not evident, excluded in the oldest apothecia, in section
to 45 µm thick, lacking photobiont cells, composed of radiating, rarely interwoven, branched hyphae
1.5 –2 µm thick, inspersed with hyaline granules that do not dissolve in KOH. Hypothecium colour-
less, to 50 µm thick, composed of short-celled hyphae to 4 µm thick, hardly interwoven. Hymenium
to 75 µm tall, colourless, ± coherent in KOH, not inspersed but overlain with an irregular layer to 8 µm
thick of yellowish brown crystals detectable under polarized light and dissolving readily in KOH.
Paraphyses 2–3 µm thick, simple, straight, rarely branched or anastomosed. Asci clavate, clearly
stalked (stalk up to 10 µm tall), 8-spored, 30 –50 × 7–15 µm (n=12), of the Lecanora-type, with ocular
chamber short and blunt. Ascospores hyaline, thick walled (1 µm), ellipsoid to oblong, occasionally
bent, simple (although old ascospores sometimes 1-septate), 10 –11.8±1.3 –15 × 4 –5.3±0.6 – 6.5 µm
(n=30) (ratio l/w= 1.69 –2.25±0.35 –2.8), irregularly arranged in the ascus. Pycnidia not seen.
Chemistry: Usnic acid and zeorin (by TLC); thallus C–, K–, PD–.
Distribution and habitat: The new species appears to be extremely rare and is known only from
the type collection, made from an old Banksia marginata tree at the margins of Melaleuca ericifo-
lia-dominated coastal woodland. It was associated with Pannaria elixii P.M.Jørg. & D.J.Galloway,
Menegazzia subpertusa P.James & D.J.Galloway, Bactropsora paludicola Kantvilas, Parmotrema
perlatum (Huds.) M.Choisy and Austroparmelina pseudorelicina (Jatta) A.Crespo, Divakar & Elix.
The woodland habitat of Lecanora helmutii represents one of the most fragile vegetation types in
Tasmania, even though this community has largely escaped the attention of conservationists and land
644 Herzogia 31 (1) Teil 2, 2018

managers. Most stands are on private land and have been extensively cleared and drained for pasture
(usually dairy). Remnant stands are usually small and fragmented, and are used as shelter for stock.
Due to browsing, there is little or no regeneration of trees and shrubs, the understorey is degraded
and weed-choked, and many have broken canopies. The result is a general “drying out” of the habitat
and a severe loss of epiphytic cryptogams. Even so, the woodlands are known to potentially support
a significant number of highly localised or rare lichens including: Bactrospora metabola (Nyl.) Egea
& Torrente (exceedingly rare in Tasmania), B. paludicola (rare, endemic to Tasmania), Malcolmiella
cinereovirens Vězda (rare), Eugeniella farinosa P.M.McCarthy & Elix (rare, endemic), Leptogium
coralloideum (Meyen & Flot.) Vain. (probably extinct in Tasmania), Punctelia subflava (Taylor) Elix
& J.Johnst. (extinct in Tasmania), Parmotrema crinitum (Ach.) M.Choisy (rare), Belonia pellucida
Coppins & Malcolm (rare), Pseudocyphellaria aurata (Ach.) Vain. (highly localised in Tasmania)
and Porina meridionalis P.M.McCarthy (rare, endemic). Lecanora helmutii represents a noteworthy
addition to this suite of highly localised species.

Figure 2: Lecanora helmutii (type). – A) Habitus. – B) Detail of the soralia. – C) Transverse section in lactophenol
cotton blue of a young apothecium, showing the excipulum of radiating hyphae. – D) Detail of an ascus showing the
typical Lecanora-type apical structure (K/I). – E) Mature ascus (K/I; DIC). – F,G) Ascospores (water, DIC). – Scales:
A,B= 0.5 mm; C= 50 µm; D,E = 5 µm; F,G= 2.5 µm.

Selected other specimens examined:

Lecanora coppinsiarum: Australia, Tasmania, Brown Mtn Rd, near Campania, 42°53'S 147°29'E, 220 m alt.,
16 August 1981, Kantvilas 741/81 & P. James [BM, HO-holotype-]]; 2 km S of Howden, 43°03'S 147°18'E, 20 m
Pérez-Ortega & Kantvilas: Lecanora helmutii, a new species from Tasmania 645

alt., 21 May 1997, Kantvilas s.n. [HO]; Three Thumbs picnic area, 42°37'S 147°53'E, 400 m alt., 11 August 2011,
Kantvilas 235/11 [HO].
Lecanora subtecta: Australia, Tasmania, Flinders Island, Yellow Beaches, 40°13'S 148°15'E, 2 m alt., 8 August
1978, Whinray 1231 p.p. [HO]; Loila Pinnacle, 41°25'S 148°11'E, 300 m alt., 1995, Duncan s.n. [HO]; c. 1.5 km N
of Little Baldy, 41°45'S 147°56'E, 720 m alt., 17 May 1999, Kantvilas 210/99 [HO]; West of Tahune Bridge, 43°06'S
146°41'E, 130 m alt., 20 July 2004, Kantvilas 217/04 [HO]. South Australia, Kangaroo Island, Mt Taylor, 35°56'S
137°03'E, 25 August 1982, Stove 1755 p.p. [AD, HO]. New South Wales, Boonoo Boonoo Falls, 28°47'51''S
152°10'00''E, 800 m alt., 4 May 2005, Elix 38306 [HO].
Lecanora symmicta aggr.: Australia, Western Australia, Sues Rd near intersection with Margaret Rd, 33°53'S
115°26'E, 45 m alt., 13 November 2011, Kantvilas 438/11 [HO, PERTH]. Queensland, Bunya Mtns, Saddle
Tree Creek, 26°52'S 151°32'E, 660 m alt., 24 November 1995, Kantvilas 179/95 [BRI, HO]; South Australia,
Kangaroo Island, Creek Bay Farm, 35°50'S, 138°05'E, 65 m alt., 12 September 2013, Kantvilas 361/13 [AD, HO].
Victoria, Brighton, Jim Willis Reserve, 37°55'23''S 144°59'14''E, 6 June 2006, Stajsic 4061 p.p. [HO]. – Germany,
Rheinland-Pfalz, south face of Mosel Valley, 50°13'40''N 7º25'36''W, 180 m alt., 25 November 2006, Pérez-
Ortega, Eichler & Cezanne [MA-Lichen 18593]. – Spain, La Rioja, Villoslada de Cameros, trail to Lake Iregua,
30TWM2052, 1800 m alt., 19 September 2004, Pérez-Ortega 539/2004 [MA-Lichen 15625].
Lecanora strobilina: Spain, Canary Islands, Tenerife, La Gomera, road from San Sebastián to Hermigua, Pinus
forest, 800 m alt., 2 August 1994, Etayo [MA-Lichen 6861]; Navarra, Arikutza reservoir, Etayo [MA-Lichen 3844].

Discussion
The new taxon is a distinctive species easily distinguished by the combination of a whitish
sorediate thallus containing usnic acid and zeorin, biatorine, yellow to cream apothecia, and
the Lecanora-type asci with hyaline ascospores.
Three other Lecanora species having biatorine apothecia with a reduced excipulum and
lacking photobiont cells and crystals are known from Australia. The main differences be-
tween those species and L. helmutii are summarized in Table 2. Lecanora symmicta (Ach.)
Ach. is recorded from Australia (Lumbsch & Elix 2004, McCarthy 2017) but, accord-
ing to Kantvilas & LaGreca (2008), specimens fitting the wide concept of L. symmic-
ta in Australia are likely to represent several different, albeit related taxa. Lecanora sym-
micta in the strict sense is characterized by biatorine apothecia, Lecanora-type asci (but
see Rodriguez Flakus & Printzen 2014) and the presence of usnic acid together with
thiophanic acid (and other xanthones) and traces of triterpenoids (Rodriguez Flakus &
Printzen 2014). Although the name L. symmictera Nyl. (Nylander 1872) has been used
occasionally for specimens of L. symmicta lacking xanthones, LaGreca & Lumbsch (2013)
pointed out that, when those compounds occur in low quantities, they are not detectable
by means of traditional chemical reactions or even TLC, and recommended that this name
should be treated as a synonym of L. symmicta. Kantvilas & LaGreca (2008) resurrected
Lecanora subtecta (Stirt.) Kantvilas & LaGreca (= Lecidea subtecta Stirt.), a species de-
scribed from Tasmania. The main differences with L. helmutii are the absence of soredia,
the orange-pink to orange-yellow apothecia (pale lemon yellow to creamy in L. helmutii),
and the remarkable presence of lemon yellow pruina in the apothecia, a rare character in
Lecanora s. lat. (Śliwa & Wetmore 2000, Śliwa et al. 2012). The thallus of L. subtecta
can become granular, but then the granules are smaller (0.1 mm diam.) than in L. helmutii.
Furthermore, the ascospores are slightly wider in L. subtecta (Table 2). This species con-
tains usnic acid and zeorin as major compounds, but minor quantities of 2,5,7-trichloro-
3-O-methylnorlichexanthone and isoarthothelin may be detected by HPLC (Kantvilas &
LaGreca 2008). Both species also differ in their ecology, with L. subtecta occurring in
eucalypt-dominated sclerophyll forest, commonly on eucalypt lignin, with most collections
coming from the cut surfaces of old eucalypt stumps.
646 Herzogia 31 (1) Teil 2, 2018

Lecanora coppinsiarum Kantvilas, recently described from Tasmania (Kantvilas 2012), also
shares several characters with L. helmutii (Table 2). However, in that species, the thallus dis-
solves entirely into granular soredia, which are usually larger (up to 50 µm in diam), and the
apothecial disc is orange-pink and covered by a coarse whitish pruina. A further, most remark-
able difference is the presence of atranorin as the only detectable secondary compound by TLC.
There are other species of the Lecanora symmicta group not known from the Australian region
that also show some resemblance to the new taxon. Lecanora confusa Almb. may also occasion-
ally have sorediate patches on its areolate thallus. It is easily distinguished from L. helmutii by the
yellowish tinge of its thallus (and soredia), its apothecia with persistent margins and fine to coarse
pruina, and the presence of xanthones together with usnic acid and, usually, zeorin (Printzen
2001). The similar Lecanora perconfusa Printzen is restricted to Baja California (Printzen 2001).
Lecanora strobilina (Sprengel) Kieffer has a similar chemistry to L. helmutii, with usnic acid and
zeorin as the major compounds, although some specimens may also contain decarboxysquamatic
and/or barbatic acids and traces of squamatic acid (LaGreca & Lumbsch 2013). This species
always has apothecia with a thalline margin, at least when young, which usually degenerates later
into yellowish-green granules (Śliwa & Wetmore 2000, Printzen 2001, LaGreca & Lumbsch
2013). Lecanora orae-frigidae R. Sant. is a characteristic species of lignicolous communities in
coastal areas in boreal and arctic areas of the Northern Hemisphere (Brodo & Vänskä 1984).
It typically has characteristic blue-grey to greyish green, well-delimited and convex soralia with
coarsely granular soredia, and contains thiophanic acid and, usually, a second xanthone, along
with usnic acid and zeorin (Brodo & Vänskä 1984, Śliwa & Wetmore 2000).
The sterile Lecanora compallens van Herk & Aptroot also contains usnic acid and zeorin; it
has a white thallus but the soredia are greenish to yellowish (Van Herk & Aptroot 1999).
Likewise, the similar L. expallens Ach. has a greenish to yellowish sorediate thallus which
contains xanthones (thiophanic acid, a xanthone similar to arthothelin, and the ‘expallens’
unknown) in addition to usnic acid and zeorin (Kukwa 2012; Tønsberg 1992).
The closest relatives of Lecanora helmutii in the ML phylogenetic analysis were L. aitema and L.
flavoleprosa (relationship not statistically supported). Lecanora flavoleprosa Tønsb. is a soredi-
ate species that is only areolate when young, with pale yellow to yellow-green soredia (30 – 65 µm
in diam). Apothecia, when present, have a dark aeruginose to dark brown colour. It contains a
range of unindentified terpenoids in addition to usnic acid and zeorin (Tønsberg 1992).
Lecanora aitema (Ach.) Hepp very much resembles a morph of L. symmicta, with darker
apothecia due to the presence of the pigment cinereorufa-green (Palice et al. 2011), although
Pérez-Ortega et al. (2010) demonstrated that it is a phylogenetically distinct taxon. Like L.
aitema, it contains usnic acid, zeorin and xanthones as major secondary metabolites (Printzen
& May 2002). The recently described Lecanora kohu Printzen et al. from Chatham Islands
(New Zealand) shows similarities with members of the Lecanora symmicta group, although it
is not phylogenetically related (Printzen et al. 2017). Apothecia in this species are immersed,
the thallus is not sorediate and it contains psoromic acid and atranorin.
Lecanora helmutii clearly belongs to the L. symmicta group (Pérez-Ortega et al. 2010) as
supported by our phylogenetic analyses, but its closest relatives still remain unknown, very
likely due to the fact that it represents the first member of this lineage from the Southern
Hemisphere to be sequenced. Much work is still required in order to clarify the taxonomy
of the group in that region, and to shed light on the biogeography of the group worldwide.
Lecanora strobilina and L. strobilinoides have been sequenced for the first time and their posi-
tion within the L. symmicta group is confirmed. Śliwa & Wetmore (2000) already pointed
Pérez-Ortega & Kantvilas: Lecanora helmutii, a new species from Tasmania 647

out the close relationship of L. strobilina with L. symmicta. LaGreca & Lumbsch (2013) even
highlighted the difficulties of separating these taxa in northeastern North America. Laundon
(2003) speculated on the relationship between L. compallens and L. strobilina, proposing the
latter to be the primary (sexual) species of the pair. Both species appear in very different
clades of the tree, but L. strobilina appears to be closely related to the sorediate L. expal-
lens. Relationships between both species were not resolved in our analyses and we agree with
Guzow-Krzemińska et al. (2017) that the genetic variability shown by L. expallens is sug-
gestive of the presence of cryptic species. Furthermore, Laundon (2003) proposed that L.
strobilinoides should be considered a geographical race of L. strobilina, since the two taxa
only differ in the number of ascospores per ascus and their size, and proposed L. strobilina
subsp. strobilinoides (Giralt & Gómez-Bolea) J.R.Laundon. In our tree both species do not
even appear as sister species, so they should be considered different taxa.

Table 2: Salient characters of Lecanora helmutii and closely related species.

metabolites

paraphyses

ascospores
hymenium
apothecial

apothecial

secondary
sorediate
pruina

disc

asci
Lecanora fine, lemon yes usnic acid, up to 30 –50  × 2–3 µm 10 –11.8 –15  ×
helmutii white yellowish (soredia zeorin 75 µm 7–15 µm 4 –5.3 – 6.5 µm
to cream up to
25 µm)
Lecanora not orange- no usnic acid, 40 –70 µm 30 – 48  × 2–2.5 µm 9 –10.5 –15  ×
symmicta pruinose brown to zeorin, ± 8 –15 µm 3 – 4 – 6 µm
pinkish thiophanic
acid and other
xanthones
Lecanora coarse, pale no (rarely usnic acid 42–50 µm 30 – 42  × 1– 8 –11.7–15 ×
subtecta lemon- orange- thallus ± (major) and ze- 9 –14 µm 1.5 µm 3 – 4.4 –5 µm
yellow pink to granular, orin (major or
yellow granules minor) (TLC);
up to 2,5,7-trichloro-
0.1 mm) 3-O-methylnor-
lichexanthone,
and isoartho-
thelin (HPLC)
Lecanora coarse, orange- yes atranorin 45 –55 µm 35 – 45  × 1·5 –2· µm 10 –11.9 –15  ×
coppinsiarum white pink (soredia 10 –15 µm 4 – 4.9 – 6 µm
up
to50 µm)

Acknowledgements
We thank Isaac GARRIDO-BENAVENT for collecting fresh specimens of Lecanora strobilinoides. SPO was sup-
ported by the grant RYC-2014-16784 from the Spanish Ministry of Economy, Industry and Competitiveness.

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Manuscript accepted: 16 April 2018.

Communicated by: Philipp Resl

Addresses of the authors

Sergio PÉREZ-ORTEGA, Real Jardín Botánico (CSIC), C/ Claudio Moyano 1, E-28014
Madrid, Spain. – E-mail: sperezortega@rjb.csic.es
Gintaras KANTVILAS, Tasmanian Herbarium, Tasmanian Museum & Art Gallery, PO Box 5058,
UTAS LPO, Sandy Bay, Tasmania 7005, Australia. – E-mail: Gintaras.Kantvilas@tmag.tas.gov.au

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